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Foraging Modes in Lacertid Lizards from Southern Africa

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Foraging Modes in Lacertid Lizards from Southern Africa Foraging modes in lacertid lizards from southern Africa William E. Cooper, Jr.1, Martin J. Whiting2 1 Department of Biology, Indiana University-Purdue University, Fort Wayne, Indiana 46805, USA e-mail: [email protected] 2 Department of Herpetology, Transvaal Museum, Paul Kruger Str., P.O. Box 413, Pretoria 0001, South Africa Abstract. Most lacertids are active foragers, but intrafamilial variation in foraging mode is greater than in most lizard families. We collected data on eight species of African lacertids to assess this variation. Both active and ambush foraging occurred within Pedioplanis and Meroles. Meroles ctenodactylus had a proportion of time moving and proportion of attacked prey detected while moving intermediate to those for actively foraging and ambushing Pedioplanis, but its number of movements per minute was exceptionally high. This species has a unique mixed foraging mode. Like active foragers, it seeks food by tongue- icking while moving and spends a high percentage of the time moving. Like ambush foragers, it searches visually for prey during pauses between movements. Our ndings con rm published data on four Kalahari lacertids. We discuss the history of foraging modes in advanced lacertids. Introduction Effective foraging behavior is a prerequisite for survival and reproduction of desert lizards, but no single foraging style is most effective for all species. Insectivorous lizards exhibit two major foraging modes: active (wide) foraging and ambush (sit-and-wait) foraging (MacArthur and Pianka, 1966; Huey and Pianka, 1981). Active foragers move through the habitat searching for prey by sight and by frequent tongue- icking to collect chemical samples (Evans, 1961; Huey and Pianka, 1981), whereas ambushers adopt ambush posts where they remain immobile while scanning visually for approaching prey (Huey and Pianka, 1981; Cooper, 1995, 1997). These foraging modes are highly stable, some entire families of lizards and even higher taxa being restricted to a single mode (Cooper, 1994, 1995, 1997; Cooper et al., 1997). However, in a few families some species are ambushers and others are active foragers (e.g., Huey and Pianka, 1981; Arnold, 1990; Cooper, 1994; Webb and Shine, 1994). Within the two major foraging modes, substantial variation occurs in the degrees and patterns of movements (Perry, 1995). c Koninklijke Brill NV, Leiden, 1999 Amphibia-Reptilia 20: 299-311 ® 300 William E. Cooper, Jr., Martin J. Whiting Differences in foraging behavior among lizards strongly in uence important aspects of lizard life history. Foraging modes are correlated with energy budgets (Anderson and Karasov, 1981, 1988), relative clutch mass during reproductive intervals (Vitt and Congdon, 1978; Huey and Pianka, 1981; Vitt and Price, 1982), defense against predators (Huey and Pianka, 1981; Vitt, 1983), and the evolution of the tongue-vomeronasal system and the associated reliance on chemosensory detection of prey (Cooper, 1994a,b, 1995, 1996a,b, 1997a,b). Despite its great importance and numerous studies of ecological and behavioral correlates of foraging mode (Huey and Pianka, 1981; Huey and Bennett, 1986; Cooper, 1994b, 1995a, 1997a), little is known about the ultimate factors that determine which foraging mode is used by a particular species. Early studies of lizards emphasized the discrete nature of the two foraging modes, but a number of authors later suggested that foraging activity may vary along some portion of a continuum from complete inactivity to continual activity rather than being restricted to two separate ends of the continuum (Magnusson et al., 1985; Pietruszka, 1986; Perry et al., 1990). There is still some disagreement, but the two foraging modes are widely recognized as distinct. McLaughlin (1989) presented evidence that lizard foraging activity does show some continuous variation within each mode, but that the range of activity does not overlap between modes. Lizard foraging activity is usually determined by measuring either or both of two variables, the proportion of the time spent moving (PTM) and number of movements per minute (MPM) (e.g., Huey and Pianka, 1981; Perry, 1995; Cooper et al., 1997). In a review of all published quantitative data on lizard foraging modes, Perry (1995) chose PTM < 0.100 as the criterion for ambush foraging. Huey and Pianka (1981) considered lizards having PTM as high as 0.143 to be ambush foragers. In addition to PTM and MPM, some investigators have measured mean speeds over the entire period of observation and during movement. In the Lacertidae, a large family of small lizards ranging throughout much of Europe, Asia, and Africa (Arnold, 1989, 1993), a large majority of species are active foragers. Among 65 species in 13 genera reviewed by Cooper (1994), only two species in two genera had been characterized as ambush foragers. Active foraging is believed to be plesiomorphic in the Lacertidae (Cooper, 1994, 1995), the relatively primitive European taxa all having been characterized as active foragers (Arnold, 1993). Ambush foraging is known only in a few species in the advanced lineage of Africa and Eurasia (Arnold and Burton, 1978; Arnold, 1990, 1993). Even in the advanced lineage, a large majority of species are active foragers. Both foraging modes are believed to exist within the subsaharan genera Pedioplanis and Meroles (Huey and Pianka, 1981; Arnold, 1990), but quantitative data are available for only six species of Kalahari lacertids, including two species of Pedioplanis differing in foraging behavior (Huey and Pianka, 1981). Here we present quantitative data on the foraging modes of eight species of lacertids representing four genera found in arid zones of South Africa and Namibia. In addition to MPM and PTM, we present information on foraging speeds and frequency of attacks on Foraging modes in African lacertids 301 prey initiated by lizards while moving and still. We compare foraging behaviors of species differing in mode in two genera, describe a mixed foraging mode in Meroles ctenodactylus, and discuss the evolution of foraging modes in advanced lacertids and possible reasons for intrageneric variation. Methods Foraging data were collected at several sites, all in South Africa except as noted. Observa- tions were conducted on Meroles ctenodactylus on 29 February and 1 March 1996 at Hon- deklipbaai (30° 18¢ S, 17° 16¢ E); M. knoxii on 29 February and 1 March 1996 at Hondeklip- baai and on 3 March 1996 at Papendorp (31° 41¢ S, 18° 13¢ E); M. reticulatus on 25 February 1996 at the foot of Dune 7 (22° 58¢ 30¢ ¢ S, 14° 40¢ 24¢ ¢ E) in Namibia near Walvis Bay; Pe- dioplanis lineoocellata in the Kalahari on 24 March at Koegechop (28° 15¢ S, 21° 5¢ E) and 26 March 1993 at Loerkop (28° 14¢ S, 20° 55¢ E); P. namaquensis in Namaqualand at Ybeep (29° 58¢ S, 17° 59¢ E) on 27 and 28 March 1993 and Wolfhoek (30° 22¢ S, 18° 12¢ E) on 29 March 1993, with single observations on 24 March at Koegechop and 26 March 1993 at Loerkop; P. undata on 11 to 14 October 1994 and 20 and 21 February 1996 at Farm Bergvellei in Namibia (19° 37¢ S, 14° 40¢ E, 20 km west of Kamanjab); Heliobolus lugubris on 24 March 1993 at Loerkop in the Kalahari and 20-21 February 1996 at Farm Bergvellei. Data were collected for a single Nucras tessellata on 28 March 1993 at Ybeep. Observations were made only on sunny days when lizards were active. We located lizards with binoculars and unaided vision while walking slowly through an area. Upon detecting a lizard, the observer stopped moving to reduce possible disturbance to the lizard. Data were recorded on microcassette tapes only for individuals that did not appear to have been disturbed. Focal animals were observed continuously for 10 min if possible, but often less if the lizard moved out of sight. Minimum acceptable observation time was 1.5 min. To ensure that data were collected only once per individual, we avoided repeated sampling in the same area. For each lizard we recorded species, locality, date, and foraging behaviors, including time moving and stationary, distance moved (within ca. 2 cm for very short movements, but ca. 0.5 m for the longest movements of ù 10 m), and feeding attempts. Postural adjustments not involving translation (e.g., turning, tail movements) were not recorded. Immobility was recorded whenever a lizard paused for one or more seconds. Shorter intervals would have been inaccurate due to competing demands of reading time, observing lizard movements, and recording both. From the movement data we calculated PTM, MPM, and two measures of foraging speed (Huey and Pianka, 1981). Moving speed was average speed while moving, i.e., distance moved divided by time spent moving. Mean speed was calculated over the entire observation interval, including time not moving. The measurements of mean speed and movement speed are rather crude due to limitations in measures of distances moved, but 302 William E. Cooper, Jr., Martin J. Whiting are more than adequate to show differences among species. We pooled data for juveniles and adults after ascertaining that there were no signi cant age differences in MPM or PTM (Mann-Whitney U tests). All of the movement measures re ect activity levels, which are sensitive to factors such as body temperature and digestion. If care is not taken to record data only for lizards that are warm enough to forage normally and have not become so full as to stop foraging, activity data may give a distorted view of foraging. Although we could not completely control effects of differing environmental temperatures, we minimized effects of temperature by collecting data only at times of day when lizards normally forage and in light and temperature conditions affording them the opportunity to maintain body temperatures at preferred levels through thermoregulation. In addition to activity, the two foraging modes have another basic difference: Ambushers initiate attacks on prey while immobile, whereas active foragers initiate attack after nding the prey while moving.
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